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241 Epidermal Characteristics of Toxic Plants for Cattle from the Salado

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241 Epidermal Characteristics of Toxic Plants for Cattle from the Salado ISSN 0373-580 X Bol. Soc. Argent. Bot. 40 (3-4): 241 - 281. 2005 Epidermal characteristics of toxic plants for cattle from the Salado river basin (Buenos Aires, Argentina) SUSANA E. FREIRE1,2, ANA M. ARAMBARRI1, NÉSTOR D. BAYÓN1, GISELA SANCHO1,2, ESTRELLA URTUBEY2, CLAUDIA MONTI1, MARÍA C. NOVOA1, and MARTA N. COLARES1 Summary: One hundred and eighty species belonging to 41 families inhabiting the Salado River Basin of the province of Buenos Aires (Argentina) were previously reported to be toxic for cattle. The purpose of this study was to provide a tool to distinguish the taxa when the plant material is desintegrated. In this way, an approach to the identification of these taxa through leaf epidermal features (anticlinal epidermal cell wall patterns, cuticular ornamentation, stomata, and hair types) is performed. A key to the 180 species as well as illustrations of diagnostic characters are given. Key words: Buenos Aires, Salado River Basin, toxic plants, anatomy, epidermal characters, stomata, hairs, Dicotyledons, Monocotyledons. Resumen: Caracteres epidérmicos de las plantas tóxicas para el ganado de la Depresión del Salado (Buenos Aires, Argentina). Las plantas tóxicas para el ganado están representadas en la Depresión del Salado (provincia de Buenos Aires, Argentina) por 180 especies pertenecien- tes a 41 familias. El objetivo del presente trabajo es determinar estos taxa a partir de material desintegrado, utilizando caracteres epidérmicos foliares (paredes anticlinales de las células epidérmicas, ornamentación de la cutícula, tipos de estomas y pelos). Se brinda una clave para la determinación de las especies e ilustraciones de los caracteres diagnósticos. Palabras clave: Buenos Aires, Depresión del Salado, anatomía, caracteres epidérmicos, estomas, pelos, Dicotiledóneas, Monocotiledóneas, plantas tóxicas. Introduction The political province of Buenos Aires is situated in central eastern Argentina. It is covered in most of its surface by a herbaceous grassy steppe, called “pampas.” The species studied are from a plain with a poorly developed drainage system. This area is known as Salado River Basin which represents approximately 80,000 km2 (Fig. 1). The principal economic activity in this area is the cattle breeding, based on natural pastures. The knowledge of the vegetation of this area is relevant for human development. Numerous floristic studies have been carried out in this area (Cabrera, 1963-1967; Vervoorst, 1967; Cabrera & Zardini, 1978; Cabrera et al., 2000). Within the diverse plant families inhabiting the Sa- lado River Basin, 180 species belonging to 41 families show toxicity for cattle (Casós, 1935; Rate- ra, 1945; Tokarnia & Dobereiner, 1982; Ragonese & Milano, 1984; Gallo, 1979; Pertusi, 1987). Epidermal traits, i.e. epicuticular wax deposition, cuticular 1 Área de Botánica, Departamento de Ciencias Biológi- cas, Facultad de Ciencias Agrarias y Forestales, UNLP, calle 60 y 119, C.C. 31, 1900 La Plata, Argentina). 2 División Plantas Vasculares, Museo de La Plata, Paseo Fig. 1. Map of Buenos Aires province showing the Salado del Bosque s/n, 1900 La Plata, Argentina. River Basin (area in gray). 241 Bol. Soc. Argent. Bot. 40 (3-4) 2005 ornamentation, epidermal cells, stomata, and hairs, have patterns used in (Table 1) was adapted from Stace proved to be an important tool in taxa delimitation in (1965): Stace’s types 1 and 2 correspond to type 1 many plant families (Metcalfe & Chalk, 1950-1979; here; Stace’s types 3 and 4 correspond to type 2 Uphof et al., 1962; Sinclair & Sharma, 1971; Lackey, here; Stace’s types 5 and 6 correspond to type 3 1978; Arambarri & Colares, 1993; Ditsch et al., 1995; here; Stace’s types 7 and 8 correspond to type 4 Barthlott et al., 1998; Stenglein et al., 2003) and also here. The anticlinal epidermal cell wall patterns are in distinguishing fragmented vegetables from feces indicated in Table 1 as: adaxial surface / abaxial and stomach contents being resistant to the digestive surface. process (Yagueddú & Cid, 1992; Pelliza et al., 1997; Stomata types were classified according Cid & Sierra, 2004). Therefore, it would be interesting to seek morphological traits that led to diagnose on causes Metcalfe & Chalk (1950, 1979) and Van Cotthem of animal mortality through feces and stomach contents. (1970), however, to establish the monocotyledons In order to achieve this goal, we apply herein histo- types, we followed Fryns-Claessens & Van morphological characters, such as leaf epidermal features. Cotthem’s (1973) classification. Leaf margin (visible in transparent leaves) was Material and Methods only used to distinguish two groups of species with glabrous leaves. Plant material studied The nomenclature follows Zuloaga et al.(1994) The study was performed using fresh leaves and Zuloaga & Morrone (1996, 1999). (collected in Buenos Aires province), and dried Epidermal characters in Poaceae were described leaves taken from herbarium specimens belonging according to the terminology of Metcalfe (1960) and to LP, LPAG, LPS, and SI (acronyms according Ellis (1979), and hair terminology follows Metcalfe Holmgren et al., 1990). The 180 taxa investigated & Chalk (1950, 1979) and Uphof (1962). For glandu- and vouchers are detailed in Appendix 1. lar hairs (Table 1) the number in brackets indicates Methods the number of head cells, and for non-glandular Fully expanded leaves were selected for the hairs the number of hair cells above the epidermis. study. Data were obtained from the central area of The observed hairs and stomata types are described the midlamina on both surfaces. For reconstitution according to Metcalfe & Chalk (1979), Ramayya of dried leaves we followed D’Ambrogio de (1962), and Uphof (1962). Argüeso (1986). Then, the material was fixed in formalin, glacial acetic acid, and 50% ethanol at a Results and Discussion 5:5:90 ratio (F.A.A.). Most of the epidermal From the 41 studied families inhabiting the Sala- microcharacters were studied by peeling and/or in do River Basin, the most representated were: samples cleared using the technique of Dizeo de Asteraceae (41 spp.), Poaceae (16 spp.), Solanaceae Strittmatter (1973). However, the replica method (14 spp.), Fabaceae (13 spp.), and Brassicaceae (10 (according to Freeman, 1984) was used in some taxa spp.). The results are presented in Table 1. where it was not possible to get epidermis by Even if stomata and indumentum types of the peeling or chloral hydrate clearing. To study the species studied are basically in agreement with those epidermal characters of the species belonging to described by Metcalfe (1960), Metcalfe & Chalk (1979, Poaceae we followed the technique of Metcalfe (1960). 1989) and Uphof (1962) for all families studied, a more The semipermanent slides were stained using saffranin detailed discussion of some families is included, in 80% ethanol and mounted in gelatine-glycerine. because they have some microcharacters that would Observations, and original drawings were made be cited for the first time and/or they present some with a light microscope, Leitz SM lux with camera luci- special traits. They are as follows: da. Measurements of stomata (length and width) and hairs were taken using a Nikon light microscope Apiaceae (6 spp. surveyed): We found equipped with an ocular micrometer. The average size anomocytic, diacytic, and paracytic stomata types; of hairs and stomata were determined based on all species showed glabrous epidermal surfaces, measurements performed on 15-20 replicates per sample. although papillae were found at midvein level in Cuticular ornamentation was cited only when it two species of Ammi. It is the first time in which was conspicuous (Table 1) diacytic stomata type has been cited for Apiaceae, The classification of anticlinal epidermal cell wall whereas anomocytic and paracytic have been 242 S. E. Freire et al., Epidermal characteristics of toxic plants for cattle Table 1. Epidermal characters of the toxic plants of Salado River Basin. Numbers in brackets by the species refer to bibliography cited below the table. aba th t b at R haabf a Cy t T ch a t C ta a a a Cy acty Cy h t Dtaa a a ch ch ac a ca y b va tat a t ca c a at L t ch ach f L tf L L t t S h ha A aa th hyb A v A a A v a a C ac at Cyc t hy y a c at c v a V ca a 243 Bol. Soc. Argent. Bot. 40 (3-4) 2005 Table 1. Continuation A c a c a avca A a abachy t ha a ata O y ta a Achy c at A b at f a A th c t a Act A t q a at q a at Baccha at B atc ata B c f a B t a Bf c B t a B a a Ca aca th C ta acactaa C t C tta Cch tyb C v a C y ab a b a Cy aaca c Gaa a a ta ca a ta ca y y a th cab a Lact ca a O aca th S c b a S c ba t att S bach bach S aaaca S tw 244 S. E. Freire et al., Epidermal characteristics of toxic plants for cattle Table 1. Continuation S v a Syb aa S a ch ch S ch ac Ta t ta Taa ac ffc a V b a c V a bca bca W a a ca a th cava a th A ch a ffc a ch a ta t a ca Ba ca a Baa Ca ab a a t Ca aa aba Ra ha a ha t Ratv Ra t R a a t t aq atc S y b at S O taa chava ta a a tt W at haca tha Sa b c a ta A t a tha yca t ta hy Sa aa ffc a S aav Ba a c aa 245 Bol.
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